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Habitat and Diet Partitioning between Shoal Bass and

Largemouth Bass in the Chipola River, Florida
a

A. P. Wheeler & Michael S. Allen

Department of Fisheries and Aquatic Sciences, University of Florida , 7922 Northwest 71st
Street, Gainesville, Florida, 32653-3071, USA
Published online: 09 Jan 2011.

To cite this article: A. P. Wheeler & Michael S. Allen (2003) Habitat and Diet Partitioning between Shoal Bass and
Largemouth Bass in the Chipola River, Florida, Transactions of the American Fisheries Society, 132:3, 438-449, DOI:
10.1577/1548-8659(2003)132<0438:HADPBS>2.0.CO;2
To link to this article: http://dx.doi.org/10.1577/1548-8659(2003)132<0438:HADPBS>2.0.CO;2

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Transactions of the American Fisheries Society 132:438449, 2003

q Copyright by the American Fisheries Society 2003

Habitat and Diet Partitioning between Shoal Bass and

Largemouth Bass in the Chipola River, Florida
A. P. WHEELER*1

AND

MICHAEL S. ALLEN

Downloaded by [UQ Library] at 07:25 13 July 2015

Department of Fisheries and Aquatic Sciences,

University of Florida,
7922 Northwest 71st Street,
Gainesville, Florida 32653-3071, USA
Abstract.We investigated the macrohabitat use, microhabitat use, and food habits of shoal bass
Micropterus cataractae and largemouth bass M. salmoides in the upper Chipola River, Florida. We
electrofished two macrohabitats (pools and shoals) during the summer (MayAugust) and fall
(SeptemberDecember) of 1999 and 2000. The ratio of shoal bass to largemouth bass differed
among macrohabitats, being highest in the shoals and lowest in the pools. Age-0 and adult (age1 and older) shoal bass were collected in areas of higher-than-average percentages of rocky substrate
in both shoals and pools. Age-0 and adult largemouth bass were associated with areas of reduced
current velocity and those with higher-than-average amounts of woody debris. Though the diets
of age-0 and adult shoal bass and largemouth bass were similar, a few differences were apparent.
Age-0 largemouth bass diets contained grass shrimp Palaemonetes spp., whereas age-0 shoal bass
diets contained mostly mayflies (order Ephemeroptera: families Baetidae and Isonychidae). Crayfish and fish were the primary food resources of adult shoal bass and adult largemouth bass, and
crayfish became more prevalent than fish in larger individuals of both species. Largemouth bass
transitioned to a crayfish-dominated diet at a smaller size than did shoal bass. Considering that
we found substantial differences in the habitat associations of these species but relatively few diet
differences, habitat partitioning may be important for the coexistence of shoal bass and largemouth
bass in rivers and streams. Future shoal bass conservation efforts should focus on maintaining a
diversity of habitats where these species coexist and on protecting relatively rare shoals.

Shoal bass Micropterus cataractae, the most recently described species of black bass Micropterus
spp., is believed to be threatened by habitat loss
throughout its limited range. Its endemic range is
the ApalachicolaChattahoocheeFlint River
drainage of Florida, Alabama, and Georgia, and
introductions have been limited to the Ocmulgee
River in Georgia (Williams and Burgess 1999).
The range of shoal bass is diminishing due to the
destruction of natural riverine habitat by impoundments (Williams and Burgess 1999). Although
shoal bass can survive and reproduce in ponds
(Smitherman and Ramsey 1972), they do not persist in impoundments (Ramsey 1975; Williams and
Burgess 1999). Thus, shoal bass have disappeared
from much of the Chattahoochee River and its tributaries in Georgia and Alabama (Williams and
Burgess 1999). Within Florida, shoal bass are restricted to a segment of the upper Chipola River
and an area below Jim Woodruff Dam on the Apalachicola River. These populations are threatened
* Corresponding author: wheelerap@brinet.com
1 Present address: North Carolina Wildlife Resources
Commission, 20830 Great Smoky Mountain Expressway, Waynesville, North Carolina 28786, USA.
Received December 7, 2001; accepted October 3, 2002

by pollution and siltation in the Chipola River

(Ogilvie 1980; Williams and Burgess 1999) and
by dredging and irregular flows in the Apalachicola River (Williams and Burgess 1999). Shoal
bass are considered a species of special concern
in Alabama (Ramsey 1976) and threatened in Florida (Gilbert 1992).
Unfortunately, a paucity of work has addressed
the biology and ecology of shoal bass. Williams
and Burgess (1999) concluded that future conservation efforts should include studies to determine
the microhabitat requirements of the species. Although shoal bass generally are thought to inhabit
shoals in rivers and streams (Ramsey 1975; Gilbert
1992; Williams and Burgess 1999), few previous
studies have attempted to verify this association.
Wright (1967) collected shoal bass and largemouth
bass M. salmoides at a 10:1 ratio from shoals and
a 3:1 ratio from pools. Hurst (1969) collected shoal
bass in both pool and riffle areas but collected
largemouth bass mainly from large, deep pools.
Shoal bass diets have been examined by few investigators but have been found to consist of fish
and crayfish (Wright 1967; Hurst 1969; Ogilvie
1980).
Shoal bass share their range with endemic largemouth bass. Largemouth bass are widely distrib-

438

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SHOAL AND LARGEMOUTH BASS HABITAT AND DIET PARTITIONING

uted, and many previous studies have examined

their biology and ecology. However, previous research has focused almost exclusively on lentic
populations (Hamilton and Powles 1983), even
though largemouth bass are common in a wide
range of lotic habitats (Jenkins and Burkhead
1993) and are often abundant in streams (Fajen
1975).
Largemouth bass are generally reported to inhabit pools and backwater areas in rivers and
streams (Trautman 1957; Wydoski and Whitney
1979) but may be habitat generalists in lotic systems. For example, Schramm and Maceina (1986)
found largemouth bass in a wide range of habitats
in the Santa Fe River, Florida. Sowa and Rabeni
(1995) found largemouth bass biomass and density
were positively correlated to maximum summer
temperature, mean depth, pool area, and pool:riffle
ratio, whereas canopy cover and gradient were
negatively correlated to largemouth bass biomass
and abundance in Missouri streams.
Largemouth bass are generally considered piscivores (Heidinger 1975) but may prey on a wide
variety of aquatic organisms (Jenkins and Burkhead 1993). Previous studies of largemouth bass
diets in lotic environments have determined that
fish were their primary prey (Scalet 1977; Davies
1981; Hamilton and Powles 1983). However,
Schramm and Maceina (1986) found that crayfish
were the primary food resource of largemouth bass
in the Santa Fe River, Florida.
Shoal bass and largemouth bass are naturally
sympatric throughout the range of shoal bass (Williams and Burgess 1999). These species are potential competitors for prey and space because they
have similar morphology and prey. Because these
fishes are naturally sympatric, they may exhibit
resource partitioning to reduce interspecific interactions and facilitate coexistence. Previous studies
of sympatric black bass populations have variously
documented differences in spatial resource use
(Sowa and Rabeni 1995; Sammons and Bettoli
1999), spatial but not food resource use (Janssen
1992; Scott and Angermeier 1998), food but not
spatial resource use (Scalet 1977), neither food nor
spatial resource use (Hubert 1977), and both spatial and food resource use (Warden and Hubert
1980; Schramm and Maceina 1986).
We investigated the habitat use and food habits
of shoal bass and largemouth bass in a lotic system.
Our objectives were to test (1) whether the ratio
of collected shoal bass to largemouth bass varied
between macrohabitats (i.e., pools and shoals), (2)
whether shoal bass or largemouth bass were as-

439

sociated with different microhabitat parameters

(i.e., current velocity, substrate, woody debris, and
depth) than the average conditions within both
pools and shoals, and (3) whether diets were similar between shoal bass and largemouth bass.
Methods
Study site.The Chipola River is a low-gradient
stream (0.17 m/km; Bass and Cox 1985) that flows
201 km south from Alabama to its confluence with
the Apalachicola River. Its watershed area is 3,124
km2, and average discharge is 42.8 m3/s (Bass and
Cox 1985). Although the Chipola River receives
some surface runoff, it is primarily spring fed (Parsons and Crittenden 1959; Bass and Cox 1985),
resulting in low turbidity (Parsons and Crittenden
1959) and relatively stable year-round water temperatures (range, 10248C; Parsons and Crittenden
1959; Bass and Cox 1985). The substrate is primarily sand and cobble, with limestone outcrops
(Parsons and Crittenden 1959). The watershed is
relatively pristine, containing some agricultural
developments and small towns (Winger et al.
1987).
The 48-km section of river between the towns
of Marianna and Clarksville, Florida, is inhabited
by shoal bass and largemouth bass. This section
of river is characterized by two primary macrohabitats, pools and shoals, with pools being the
dominant macrohabitat. Pools are characteristically narrow (mean width, 31 m), long (.1 km),
deep (mean, .2 m), and slow flowing. Shoals are
shorter (,200 m), wider (mean, 66 m), shallower
(mean depth, ,1 m), and have higher current velocities than the pools. Shoals also contain beds
of eelgrass Vallisneria americana, whereas pools
are devoid of aquatic macrophytes. We selected
three pools and three shoals for sampling. The
three shoals were locations where previous studies
collected shoal bass, and the pools were adjacent
to the shoals. All study units were located within
a 15-river-kilometer (rkm) section.
Fish collection.Fish and microhabitat sampling were conducted in summer (MayAugust)
and fall (SeptemberDecember) of 1999 and 2000.
Fish were collected by boat electrofishing with
DC. Habitat differences between pools and shoals
required different electrofishing techniques for effective fish collection. Shoals were electrofished
by moving upstream in such a way that all areas
deep enough to navigate were exposed to the electric field. Due to the presence of shallow areas
(,0.25 m), the entire area of the shoals could not
be navigated; however, many areas too shallow to

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440

WHEELER AND ALLEN

navigate were within reach of the cathode probe.

Pools were electrofished by drifting longitudinally
down the habitat unit while moving laterally between the right bank, left bank, and the center of
the channel. Units were sampled as many as three
times in each season but always with at least 48
h between samples. Because pools are the primary
macrohabitat in the upper Chipola River, we
weighted our sampling effort in favor of pools.
Areas where shoal bass or largemouth bass were
collected were marked with a weighted buoy for
microhabitat measurements. The buoys were color
coded to distinguish between age-0 fish and those
older than age 0 (hereafter referred to as adults)
and between shoal bass versus largemouth bass.
Ages of fish that appeared to be age 0 in the field
were verified by examining sagittal otoliths in the
laboratory. Ad hoc sampling in areas adjacent to
the three shoals and three pools was used to supplement fish collections for diet analysis.
Macrohabitat analysis.Because habitat differences between pools and shoals required different
electrofishing techniques, we could not use electrofishing catch per effort to compare the relative
abundances of shoal bass and largemouth bass between macrohabitats. We tested whether the proportion of shoal bass to largemouth bass differed
among macrohabitats with a weighted-leastsquares analysis of proportions (WLSAP; Statistical Analysis System [SAS] Catmod procedure;
SAS 1994; Stokes et al. 2000). The WLSAP tested
whether the proportion of shoal bass to largemouth
bass differed across macrohabitat types, seasons,
and years.
Microhabitat characterization and analysis.
We characterized the microhabitat associations of
age-0 and adult shoal bass and largemouth bass by
measuring or estimating physical habitat at the location where individuals were collected. Microhabitat parameters were measured at an area designated longitudinally (i.e., upstreamdownstream
axis) as 10 m wide, centered at the point the individual was captured, and latitudinally as the left,
right or center third of the stream. The area contained in each interval was variable, dependent on
stream width.
We characterized microhabitat conditions in the
pools and shoals using four permanent transects
in each habitat unit. Permanent transects were perpendicular to the channel and approximately evenly spaced through each unit. A tree on the bank
adjacent to the transect was marked with paint,
allowing the same fixed transects to be used
throughout the study. During each sampling event

(i.e., summer and fall of 1999 and 2000), microhabitat characteristics were recorded from three
intervals equally spaced across each transect (e.g.,
two near the bank and one midchannel). The longitudinal width (i.e., upstreamdownstream axis)
of the intervals was defined as 10 m on either side
of the transect. The area contained in each interval
was variable, dependant on stream width. Thus,
the intervals of the fixed transects were statistically
comparable to the areas where fish were collected.
Microhabitat and cover variables were measured
or estimated for each interval of the fixed transects
and for the areas where individual shoal bass or
largemouth bass were collected. Mean depth was
estimated to the nearest 5 cm with a depth rod.
Mean current velocity was measured at 60% of
depth, at a point representative of the interval
(judged visually), with a Marsh-McBirney model
201M portable water current meter. Substrate was
recorded as the visually estimated percentage of
silt (,0.062 mm), sand (.0.0622 mm), gravel
(.216 mm), pebble (.1664 mm), cobble (.64
256 mm), boulder (.256 mm), and flat bedrock.
The total percentage of the substrate classified as
cobble and boulder was summed and used to indicate rocky substrate particles. Rocky substrate
percentage was used in the analysis because it provided an easily interpretable index of substrate
coarseness and because there is a reported association of other black basses with these substrates
(Leonard and Orth 1988; Todd and Rabeni 1989;
Lobb and Orth 1991). The percent area covered
by eelgrass was also visually estimated. Individual
pieces of woody debris were visually counted in
four categories (IIV) based on length and diameter, as done by Dolloff et al. (1993). The four
categories we used were: (I) 15 m in length and
510 cm in diameter, (II) 15 m in length and over
10 cm in diameter, (III) over 5 m in length and 5
10 cm in diameter, and (IV) over 5 m in length
and over 10 cm in diameter. In the case of fallen
trees, branches connected to a larger trunk were
categorized and enumerated individually. From
these counts and categories, a woody debris index
(WDI) was calculated as
WDI 5 (Wood I) 1 [2 3 (Wood II)]
1 [3 3 (Wood III)]
1 [4 3 (Wood IV)],

(1)

where Wood IIV represent numbers of woody debris of each size category in the interval. This
index was weighted so that larger size-classes of

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SHOAL AND LARGEMOUTH BASS HABITAT AND DIET PARTITIONING

debris received an arbitrarily higher value. Larger

values of WDI are used to indicate greater amounts
of woody debris in the interval. Percent area coverage of eelgrass was estimated visually in each
interval. The first author measured or estimated all
microhabitat parameters except mean current velocity throughout the study.
We validated the visual estimation of substrate
composition. Substrate composition of 15 approximately 25-m2, shallow (depth, ,1 m) locations in
a shoal was visually estimated as previously described. Locations were selected to reflect as much
variation in substrate particle composition as possible. After visually estimating the substrate at a
location, we randomly tossed a 0.5-m2 ring, divided by string into 16 equal sections, into the
location 10 times. The dominant substrate particle
category was recorded in each of the sections for
every toss. For each location, the percentage of
the 160 sections (16 sections 3 10 tosses) dominated by rocky substrate was compared to the visual estimation of percent rocky substrate. The areas used to verify the visual estimation of substrate
particles contained substrates ranging from sand
to boulder. There was a significant linear correlation (R 5 0.94, P , 0.001) between the visual
estimation of rocky substrate and the observed
proportion of sections dominated by rocky substrate. Thus, the visual estimation appeared to be
an adequate index of substrate composition.
We used a multivariate analysis of variance
(MANOVA) to test the null hypothesis that microhabitat variables collectively were similar between the permanent transects (average conditions
of the macrohabitat) and the areas where individuals were collected. The analysis was repeated for
age-0 and adult individuals of both species. The
MANOVAs were conducted for individuals in
pools and shoals (eight analyses total) based on
the microhabitat observations (i.e., intervals) from
the fixed transects and the microhabitat data from
areas where individuals were collected. Current
velocity, depth, percent rocky substrate, and WDI
score were used as response variables, and year
(1999, 2000), season (summer, fall), species presence (e.g., at least one individual was or was not
collected), and the interaction between species
presence and season were used as predictor variables. In addition, eelgrass abundance was used as
a response variable for the shoals. Data were log10
transformed as necessary to help meet the assumption of a multivariate normal distribution
(Zar 1988). We used Wilks lambda (SAS 1994)
to test the null hypothesis that year, season, species

presence, and the species presence 3 season interaction had no effect on the mean microhabitat
observations. In cases of an overall significant
MANOVA for the effect of species presence, oneway analyses of variance (ANOVA) were used to
determine microhabitat parameters that contributed to the significant difference between the areas
where individuals were collected and the permanent transects.
Diet collection and analysis.Transparent
acrylic tubes (Van Den Avyle and Roussel 1980)
were used to remove the stomach contents of all
shoal bass and largemouth bass larger than 200
mm total length. A flexible claw retriever (Dimond
1985) assisted in the removal of large items and
crayfish. The removed stomach contents were
placed on ice and returned to the laboratory, where
the items were enumerated, identified, and
weighed. Shoal bass and largemouth bass 200 mm
or smaller were placed on ice and returned to the
laboratory, where they were dissected and their
stomach contents removed, enumerated, identified,
and weighed to the nearest 0.001 g. When possible,
fish remains were identified to genus (based on
Oates et al. 1993) and insects to order.
Diet was quantified for each species in terms of
percent by weight. Piankas (1973) index of diet
overlap (O) was used to index the similarity of the
diets of the two species. Piankas Index is defined
as:
O5

O p p @1O p 3 O p 2
2i 1i

2i

1i

0.5

(2)

where O is the overlap in resources between species 1 and 2, p1i is the weight of resource category
i found in species 1, and p2i is the weight of resource category i found in species 2. The resource
categories were fish, crayfish, insects, other invertebrates, and other prey items. Overlap (i.e.,
similarity) values range from 0 (none) to 1 (complete). EcoSim software (Gotelli and Entsminger
1997) was used to generate null models and to
calculate the probability of observed diet overlap
occurring by chance. Driscoll and Miranda (1999)
used EcoSim software to evaluate diet overlap
among age-classes of yellow bass Morone mississippiensis in Mississippi River oxbow lakes. A randomization algorithm (RA2, Gotelli and Entsminger 1997) was selected that generated null models
for each species based on only those prey items
observed in the diets.
Ontogenetic diet shifts were evaluated with logistic regression. Logistic regression models the
probability of a binary response based on contin-

442

WHEELER AND ALLEN

and b1b3 are the logistic regression coefficients.

The estimate of the logit(p) was used to obtain the
predicted probability of a diet being crayfish dominated, that is,
p 5 elogit(p)/(1 1 elogit(p)).

(4)

Walds chi-square statistic (SAS 1994) was used

to test the significance of the individual model
terms.

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Results
The total amount of electrofishing time was 88.6
h, accumulated from pool (51.5 h), shoal (26.1 h),
and ad hoc (10.9 h) collections. A total of 105
adult shoal bass, 316 adult largemouth bass, 288
age-0 shoal bass, and 125 age-0 largemouth bass
were collected during this study. Shoal bass ranged
from 70 to 480 mm TL, whereas largemouth bass
ranged from 60 to 560 mm TL (Figure 1).
The study area generally exhibited slightly
lower-than-average monthly discharges for both
study years, and average monthly flows were higher in 1999 than 2000. Mean depth and current velocity were greater in 1999 than 2000 in both pools
and shoals (Table 1) due to higher rainfall in 1999.
The percent rocky substrate was relatively consistent throughout the study. Eelgrass was more abundant in 2000 (Table 1).

FIGURE 1.Length-frequency histogram showing size

(TL; 20-mm groups) and number of shoal and largemouth bass collected during 1999 and 2000 combined.

uous and/or categorical predictor variables (SAS

procedure Logistic; SAS 1994; Stokes et al. 2000).
In this application, the binary response variable
was fish or crayfish remains dominating (by
weight) the diet of shoal bass or largemouth bass.
The linear model used was

Macrohabitat Results

logit( p) 5 a 1 b1 3 TL 1 b2 3 species
1 b3 3 (species 3 TL),

Age-0 shoal bass and largemouth bass were collected from both pools and shoals (Table 2). However, the WLSAP detected differences in the ratio
of age-0 shoal bass to age-0 largemouth bass in
pools and shoals. We collected too few age-0 individuals in 1999 to test for differences between
the two sample years (Table 2). Therefore, the
years were pooled for this analysis. Macrohabitat
type was the sole significant predictor of the ratio
of age-0 shoal bass to age-0 largemouth bass

(3)

where logit(p) is the logistic probability of crayfish

(versus fish) dominating the diet of either shoal
bass or largemouth bass, a is the intercept value,
TL is the total length (mm) of each species, species
is the main effect of the categorical variable species type (shoal bass or largemouth bass), species
3 TL is the interaction between species and TL,

TABLE 1.Mean 6 2 SEs observations for depth, current velocity, percentage rocky substrate, woody debris index
(WDI), and eelgrass (% coverage) in the fixed transects of the pools (N 5 36) and shoals (N 5 36) during summer
(MayAugust) and fall (SeptemberDecember), 1999 and 2000, in the Chipola River.
Habitat

Year

Season

Pool

1999

Summer
Fall
Summer
Fall
Summer
Fall
Summer
Fall

2000
Shoal

1999
2000

Depth (cm)
195
173
158
154
105
69
63
67

6
6
6
6
6
6
6
6

11.6
9.6
9.5
9.8
9.5
6.1
5.7
6.3

Velocity
(cm/s)
39
26
15
13
51
31
27
19

6
6
6
6
6
6
6
6

2.1
2.0
1.6
1.4
5.3
2.6
3.2
2.1

Rocky
substrate
23
28
29
33
29
43
42
42

6
6
6
6
6
6
6
6

4.2
3.8
3.7
4.7
3.3
2.6
3.0
3.2

WDI
3
2
2
,1
2
2
2
1

6
6
6
6
6
6
6
6

1.3
0.8
0.5
0.3
0.6
0.7
0.6
0.3

Eelgrass
,1
,1
,1
10
9
26
1
22

6
6
6
6
6
6
6
6

0.3
0.1
0.1
4.1
3.1
4.8
0.8
5.2

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SHOAL AND LARGEMOUTH BASS HABITAT AND DIET PARTITIONING

TABLE 2.Number of fish collected from pool and

shoal sample sites in the Chipola River as well as from ad
hoc sampling of nearby areas that was used to supplement
fish collections for diet contents.
Sample
Species and age-class

Pool

Shoal

Ad hoc

Age-0 shoal bass

Adult shoal bass
Age-0 largemouth bass
Adult largemouth bass

118
83
82
252

137
26
20
34

33
9
23
33

TABLE 3.Results of MANOVA analyses testing for

differences between the mean habitat observations in intervals where fish were collected (presence) versus the
fixed intervals. In addition, effects of season (summer or
fall), year (1999 or 2000), and the interaction between
presence and season were included in the MANOVA. Because age-0 fish of both species did not fully recruit to the
gear until fall sampling, fall habitat observations were
used in the MANOVA analyses for age-0 shoal and largemouth bass.
Wilks
lambda

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Effect

(x2 5 41.77, df 5 1, P , 0.0001). The ratio of

age-0 shoal bass to age-0 largemouth bass was
significantly higher in shoals (6.9:1) than in pools
(1.4:1).
Adult shoal bass and largemouth bass were collected from both pools and shoals (Table 2). However, the WLSAP also detected differences in the
ratio of adult shoal bass to adult largemouth bass
in these macrohabitats. The ratio of adult shoal
bass to adult largemouth bass was predicted by
year (x2 5 10.88, df 5 1, P 5 0.0010), macrohabitat (x2 5 8.08, df 5 1, P 5 0.0945), and season
(x2 5 21.41, df 5 1, P , 0.0001). The main effect
of year was significant because the ratio of adult
shoal bass to adult largemouth bass was lower in
2000 (1:4.1) than in 1999 (1:3.51). The significant
main effect of season resulted from the collection
of a higher ratio of adult shoal bass to adult largemouth bass in the summer sampling (1:2.0) than
in fall sampling (1:4.3). The ratio of adult shoal
bass to adult largemouth bass was greater in the
shoals (1:1.3) than in the pools (1:3.1).
Microhabitat Results
All MANOVA analyses detected significant differences between the microhabitats where age-0
and adult shoal bass and largemouth bass were
collected and the average conditions of both pools
and shoals (Table 3). The presence 3 season interaction was never significant (all P . 0.06), indicating that generally these species did not change
microhabitat associations seasonally. The main effect of presence was always significant (all P ,
0.008), indicating that both species were associated with microhabitat parameters that differed
from the mean microhabitat parameters in pools
and shoals (Table 3).
Age-0 and adult shoal bass showed similar
trends in microhabitat associations. Age-0 and
adult shoal bass were associated with deeper-thanaverage areas in shoals (one-way ANOVA: F1,149
5 18.1, P , 0.001; F1,155 5 6.1, P 5 0.015) and

df

P-value

Presence
Year

Age-0 shoal bass in pools

0.8531
6.6
4, 154
0.8270
8.1
4, 154

,0.001
,0.001

Presence
Year

Age-0 shoal bass in shoals

0.8608
4.7
5, 145
0.8785
4.0
5, 145

,0.001
0.002

Presence
Year

Age-0 largemouth bass in pools

0.8005
7.1
4, 114
0.6588
14.8
4, 114

,0.001
,0.001

Presence
Year

Age-0 largemouth bass in shoals

0.8203
3.4
5, 77
0.8725
2.3
5, 77

0.008
0.058

Presence
Season
Presence 3 season
Year

Adult shoal bass in pools

0.9397
3.3
4,
0.9340
3.1
4,
0.9850
0.7
4,
0.6229
26.5
4,

175
175
175
175

0.012
0.017
0.617
,0.001

Presence
Season
Presence 3 season
Year

Adult shoal bass

0.8968
0.9400
0.9703
0.7625

151
151
151
151

0.005
0.093
0.468
,0.001

Adult largemouth bass in pools

Presence
0.7416
20.0
4, 229
Season
0.9299
4.3
4, 229
Presence 3 season
0.9773
1.3
4, 229
Year
0.7030
24.2
4, 229

,0.001
0.002
0.260
,0.001

Adult largemouth bass in shoals

Presence
0.7068
13.7
5, 165
Season
0.9553
1.5
5, 165
Presence 3 season
0.9403
2.1
5, 165
Year
0.7867
9.0
5, 165

,0.001
0.179
0.069
,0.001

in shoals
3.5
5,
1.9
5,
0.9
5,
9.4
5,

shallower-than-average areas in pools (F1,157 5

12.4, P , 0.001; F1,178 5 5.5, P 5 0.020). Age-0
and adult shoal bass were collected in areas of
higher-than-average rocky substrate in shoals
(F1,149 5 4.9, P 5 0.029; F1,155 5 9.9, P 5 0.002)
and pools (F1,149 5 16.3, P , 0.001; F1,178 5 10.3,
P 5 0.002). Adult shoal bass were found at lowerthan-average eelgrass coverage in the shoals (F1,155
5 5.2, P 5 0.024).
Age-0 and adult largemouth bass also showed
similar trends in microhabitat associations. Age-0
and adult largemouth bass were associated with

444

WHEELER AND ALLEN

deeper-than-average areas in the shoals (one-way

ANOVA: F1,81 5 8.5, P 5 0.005; F1,169 5 24.9,
P , 0.001) and shallower-than-average areas in
the pools (F1,117 5 8.8, P 5 0.004; F1,232 5 5.8,
P 5 0.017). Age-0 and adult largemouth bass were
collected from lower-than-average current velocities in pools (F1,117 5 3.8, P 5 0.054; F1,232 5
5.3, P 5 0.022). In both pools and shoals, adult
largemouth bass were associated with areas of
higher-than-average WDI scores (F1,232 5 67.5,
P , 0.001; F1,169 5 22.0, P 5 0.001). Adult largemouth bass were found at lower-than-average coverage of eelgrass in the shoals (F1,169 5 16.1, P ,
0.001).

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Diet Analyses
Stomach contents were examined for 288 (77
empty) age-0 shoal bass and 125 (25 empty) age0 largemouth bass. All diets collected in 1999 were
weighed, but because higher numbers of age-0
shoal bass and age-0 largemouth bass were collected in 2000, the age-0 diets were subsampled.
Individuals collected during 2000 were randomly
subsampled until 10 (nonempty) individuals per
length group were examined. There was a higher
overlap in the diets of age-0 shoal bass and age0 largemouth bass than would be expected to occur
by chance alone (O 5 0.88, P 5 0.054), indicating
that their diets were generally similar. The diets
of most age-0 shoal bass and age-0 largemouth
bass were dominated by unidentifiable fish, and
both species exhibited increasing prevalence of
fish in the diets with increasing TL (Figure 2).
However, the high diet overlap values may exaggerate diet similarity due to the prevalence of
relatively heavy fish remains. Differences in diets
were apparent when prey items were compared by
size-classes, especially for individuals less than
150 mm TL (Figure 2). Age-0 shoal bass diets were
dominated by mayflies (order Ephemeroptera:
families Baetidae and Isonychidae), whereas age0 largemouth bass showed a higher prevalence of
grass shrimp Palaemonetes spp. in their diets (Figure 2). Diets consisting of over 200 individual
mayflies were common in age-0 shoal bass, but
this phenomenon was not observed in age-0 largemouth bass.
Stomach contents of 118 (35 empty) adult shoal
bass and 319 (124 empty) adult largemouth bass
were examined. There was a higher overlap in the
diets of adults of both species than would be expected to occur by chance alone (O 5 0.86, P 5
0.073). Crayfish and fish dominated the adult diets
of both species (Figure 2). Although the majority

of fish remains were unidentifiable, darters Percina

spp. and madtoms Noturus spp. were most common in the diets of adult shoal bass, whereas darters and sunfishes Lepomis spp. were most prevalent
in the diets of adult largemouth bass.
The logistic regression analysis revealed the occurrence of an ontogenetic shift in the diets of adult
shoal bass and adult largemouth bass. Crayfish became increasingly more dominant than fish in the
diets of large individuals of both species (main
effect of TL x2 5 46.9, P , 0.001, Figure 3). The
diet shift is modeled by:
logit( p) 5 23.2108 1 0.0113 3 TL
2 0.7767 3 (species)
1 0.00541 3 (species 3 TL)

where species is equal to 21 for shoal bass and

11 for largemouth bass. Largemouth bass made
the transition to a crayfish-dominated diet faster
and at a smaller TL than did shoal bass (species
3 TL interaction x2 5 10.8, P 5 0.001, Figure 3).
For example, the logistic regression predicted that
a 239-mm TL largemouth bass would have an
equal probability of displaying a crayfish-dominated or fish-dominated diet, whereas for shoal
bass, such diets did not reach equal probability
until 413 mm TL.
Discussion
Age-0 and adult shoal bass and largemouth bass
were collected from both pools and shoals, indicating that both species were somewhat generalistic in their macrohabitat associations. The upper
Chipola River contains primarily pool habitat, and
shoal habitat makes up a relatively minor proportion of the area. Thus, we invested greater effort
in sampling pools, and we collected more shoal
bass (except age-0 fish) and largemouth bass from
the pools. Physical differences in pools and shoals
prevented us from using electrofishing catch per
effort to assess differences in the abundance of
these species across macrohabitats. However, we
detected differences in the ratio of one species to
another in the different macrohabitats, which indicated habitat differences between these two species. The ratio of shoal bass to largemouth bass
was highest in the shoals and lowest in the pools,
indicating that shoal bass represented a significantly greater percentage of the black bass assemblage in shoals. This trend was consistent for both
age-0 and adult fish and consistent with previous
studies (Wright 1967; Hurst 1969).

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SHOAL AND LARGEMOUTH BASS HABITAT AND DIET PARTITIONING

FIGURE 2.Percent by weight of prey items in the diets of shoal and largemouth bass collected during 1999 and
2000 combined. The number of stomachs containing prey items is listed over the respective length category column.
Diets of individuals smaller than 200 mm TL were subsampled in 2000.

Macrohabitat associations of age-0 individuals

may have been influenced by differing nesting
preferences of shoal bass and largemouth bass.
Previous studies have observed shoal bass nests
in a shoal (Wright 1967) and just upstream of a
riffle (Hurst 1969), whereas largemouth bass in
lotic systems are believed to nest in pools (Jenkins
and Burkhead 1993). Given the reported nesting
preferences of shoal bass and the high proportion

of age-0 shoal bass in shoals, these areas may represent spawning habitat for adults, nursery habitat
for shoal bass, or both. However, we did not document spawning habitat or movement of either
species, which warrants further investigation.
Age-0 and adult individuals of both species
showed differences in habitat use from the mean
microhabitat observations in the pools and shoals.
Age-0 and adult individuals of both species were

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446

WHEELER AND ALLEN

FIGURE 3.The relationship between TL (mm) and

the probability of an individual largemouth bass or shoal
bass exhibiting a crayfish-dominated diet (by weight).
Results of the logistic regression analysis are shown (see
text).

associated with deeper-than-average areas in the

shoals and shallower-than-average areas in the
pools. These observations likely result from a gear
bias, because our electrofishing equipment was not
able to reach all the shallow areas in the shoals
and was likely ineffective in sampling the deepest
areas in the pools (.2 m). Therefore, the habitat
associations we identified probably exclude shallow areas in shoals and deep sections of the pools.
Shoal bass were associated with areas of higherthan-average percentages of rocky substrate,
whereas largemouth bass were not. Age-0 shoal
bass were associated with higher-than-average percentages of rocky substrate in pools, and adult
shoal bass were associated with higher-thanaverage percentages of rocky substrate in both
pools and shoals. Cobble and boulder substrate is
commonly considered an important habitat type
for smallmouth bass M. dolomieu. Age-0 smallmouth bass commonly associate with cobble substrate (Leonard and Orth 1988; Livingstone and
Rabeni 1991), and adults are known to use boulders as cover in areas that are surrounded with
other substrates, such as cobble (Leonard and Orth
1988; Todd and Rabeni 1989; Lobb and Orth
1991). Our results suggest similar habitat associations for shoal bass.
Largemouth bass are believed to inhabit pools
and backwater areas in lotic situations (Trautman
1957; Wydoski and Whitney 1979). We found evidence to support this presumption. The ratio of
shoal bass to largemouth bass was lowest in the

pools, and age-0 and adult largemouth bass were

collected in lower-than-average current velocities
in pools. Shoal bass were not associated with areas
of lower-than-average current velocity in either
pools or shoals. Miller (1975) reported that largemouth bass inhabited pools and backwater areas
when they occurred in streams with smallmouth
bass, spotted bass M. punctulatus, and Suwannee
bass M. notius. Sowa and Rabeni (1995) found
largemouth bass were most abundant in streams
with high pool:riffle ratios. In addition, Schramm
and Maceina (1986) found intermediate-sized
(149299 mm TL) largemouth bass were most
abundant in areas of relatively low current velocity
(325 cm/s) in the Santa Fe River, Florida. Thus,
pools may be the preferred habitat of largemouth
bass in lotic systems. However, Schramm and Maceina (1986) collected large (.300 mm TL) largemouth bass primarily from a turbulent area with
high current velocity (3092 cm/s) and bedrock
substrates. Similarly, we collected some largemouth bass in shoals, suggesting that largemouth
bass may be more general in their stream habitat
associations than previously believed.
Adult largemouth bass were associated with
higher-than-average WDI scores in both the pools
and shoals. Age-0 largemouth bass were also associated with higher-than-average WDI scores in
the pools. The importance of woody debris in
warmwater streams is well documented. Woody
debris provides increased invertebrate production
(Benke et al. 1985), protection from strong current
velocity (Todd and Rabeni 1989), and camouflage
from predators or prey (Angermeier and Karr
1984). However, Lehtinen et al. (1997) found no
relationship between largemouth bass habitat associations and woody debris in the upper Mississippi River, and Sowa and Rabeni (1995) found
no relationships between woody debris and largemouth bass density or abundance in Missouri
streams. We found the mean WDI for areas where
largemouth bass were collected was higher than
the average values in both pools and shoals. Largemouth bass may use woody debris as structure
from which to ambush crayfish, as shelter from
current, or they may simply be responding to an
innate association with this type of structure.
Woody debris index scores for microhabitats
where shoal bass were collected did not differ from
average WDI scores in the pools or shoals.
Generally, we found similar food habits between
shoal bass and largemouth bass, with high (O .
0.8) diet overlap. The estimate of diet overlap may
be inflated since only five prey categories (fish,

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SHOAL AND LARGEMOUTH BASS HABITAT AND DIET PARTITIONING

crayfish, insects, other invertebrates, and other)

were used in the analysis. Some differences in food
habits were apparent when food taxa were separated into smaller categories. For example, fish
were the primary prey (by weight) of age-0 largemouth bass over 80 mm TL and age-0 shoal bass
over 120 mm TL. Prior to the onset of piscivory,
age-0 shoal bass primarily consumed baetid mayfly larvae. Age-0 largemouth bass rarely consumed
mayflies but frequently consumed grass shrimp,
which were almost absent in the diets of age-0
shoal bass. The consumption of grass shrimp and
the overall diet of age-0 largemouth bass were similar to the previous observations of Davies (1981)
and Schramm and Maceina (1986). Minor differences in the diets of age-0 shoal bass and age-0
largemouth bass may have resulted from differences in prey availabilities in the microhabitats
and macrohabitats occupied by the two species.
Because of low sample size (Figure 2) for some
size categories of shoal bass and largemouth bass,
we were unable to analyze diet differences between the pools and shoals. Nevertheless, the prey
consumed by both species were generally similar
in this study.
Fish and crayfish were the primary prey of adult
shoal bass and adult largemouth bass, and the prevalence of crayfish in their diets increased with fish
size. However, this transition to a crayfishdominated diet occurred at a smaller size for largemouth bass than for shoal bass. Adult largemouth
bass larger than 239 mm TL preyed almost entirely
on crayfish, whereas shoal bass diets were not as
exclusive. Roell and Orth (1993) observed a similar trend in the diets of smallmouth bass, and
Schramm and Maceina (1986) documented this
phenomenon in the diets of largemouth bass and
Suwannee bass in a Florida River. This study
agrees with previous studies, showing that fish
(Wright 1967; Ogilvie 1980) and crayfish (Hurst
1969) are the primary food resources of shoal bass.
Similar species in sympatric situations frequently exhibit resource partitioning, presumably to facilitate coexistence (Hardin 1960). We found substantial differences in the habitat associations of
shoal bass and largemouth bass in this study, but
comparatively few differences in food habits. Previous studies have shown spatial resource partitioning is more common than food resource partitioning among similar species (Schoener 1974),
congenerics (reviewed in Ross 1986), and centrarchids (Werner and Hall 1977; George and Hadley 1979; Janssen 1992; Scott and Angermeier
1998). Our study also suggests that habitat parti-

447

tioning may be more important than diet differences for the coexistence of shoal bass and largemouth bass in the Chipola River, Florida.
Maintaining shoal and pool habitats, as well as
a diversity of microhabitats such as rocky substratum and woody debris, may be important for
facilitating the coexistence of shoal bass and largemouth bass in streams and rivers. Shoal bass are
consistently extirpated from impounded sections
of rivers (Ramsey 1975; Williams and Burgess
1999), whereas largemouth bass persist. Given that
shoal bass can reproduce in ponds, Ramsey (1975)
suggested that competition with lake-dwelling
fishes eliminates shoal bass from impoundments.
Our results suggest that shoal bass and largemouth
bass partitioned resources spatially in the Chipola
River. Thus, future shoal bass conservation efforts
should focus on maintaining or enhancing stream
habitat diversity and on protecting the relatively
uncommon shoals.
Acknowledgments
We thank T. Bonvechio, M. Bledsoe, K. Dockendorf, C. Hanson, K. Henry, J. Howard, B. Pine,
and K. Tugend for assistance with field work.
Comments by P. Angermeier, P. Bettoli, T. Frazer,
D. Murie, W. Neal, and T. Newcomb improved this
manuscript. This manuscript represents publication number R-09272 of the Florida Agricultural
Experiment Station.
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